ECOLOGY AND DISTRIBUTION OF THE FLORIDA BOG FROG AND FLATWOODS SALAMANDER ON EGLIN AIR FORCE BASE David C. Bishop Dissertation submitted to the faculty of the Virginia Polytechnic Institute and State University in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Fisheries and Wildlife Sciences Carola Haas, Chair Mike Aust Jim Fraser Jim Parkhurst Jeff Walters June 9, 2005 Blacksburg, Virginia Keywords: amphibian, behavior, call, conservation, fire, tadpole Copyright 2005, David C. Bishop Ecology and distribution of the Florida bog frog and flatwoods salamander on Eglin Air Force Base David C. Bishop ABSTRACT I studied the ecology and distribution of the Florida bog frog (Rana okaloosae) and flatwoods salamander (Ambystoma cingulatum) on Eglin Air Force Base in northwest Florida. I report data on the breeding ecology, population dynamics, home ranges, microhabitat, and distribution of the endemic bog frog and make comparisons to its closest relative, the bronze frog (Rana clamitans clamitans). Bog and bronze frogs occur in the same habitats and are suspected to hybridize. I investigated the potential for auditory and visual mate recognition errors between R. okaloosae and R. clamitans. I also described the vocal repertoire of the bog frog and observed the response of resident males to the playback of unfamiliar conspecific and heterospecific (R. clamitans) calls. The advertisement calls of bog frogs vary among individuals, and individual voice recognition is possible. I exposed tadpoles of bog frogs, bronze frogs, and leopard frogs (R. sphenocephala) to chemical cues from 2 predators, the banded water snake (Colubridae: Nerodia fasciata) and the red fin pickerel (Esocidae: Esox americanus) to evaluate whether swimming behavior or habitat selection differed among tadpole species. The time spent swimming differed among tadpole species and predator treatments, differences which potentially could affect growth rates, survivorship rates, and distribution patterns. Lastly, I discuss the relationship between fire and the federally–threatened flatwoods salamander (Ambystoma cingulatum). I compared 13 breeding wetlands with different fire histories in addition to burned and unburned halves of a partially–burned wetland. In general, areas that burned more recently had more open canopies, higher dissolved oxygen concentrations, higher water temperatures, more understory vegetation, and were shallower than unburned areas. Rangewide surveys indicate that prescribed fires typically are applied in winter and early spring in areas inhabited by flatwoods salamanders. Based on what is known about the natural history of the species, the historical burning regime of the longleaf ecosystem, and the effects of fires on ephemeral wetlands, I suggest that land mangers diversify their fire–management strategy to increase the likelihood of burning the breeding wetlands of flatwoods salamanders. iii ACKNOWLEDGEMENTS This research was supported by a grant from the natural resources branch (Jackson Guard) at Eglin Air Force Base and administered through the USGS Biological Resources Division, Virginia Cooperative Fish and Wildlife Research Unit of Virginia Tech. Throughout this document, I use the singular pronoun to maintain consistency; however, I had significant help from others. I thank my advisor, Carola Haas, for her help with all aspects of this project. She allowed me to conduct my research in my own direction and always was available when I needed guidance. She also fed me on occasion and showed me that she picks a pretty– good banjo. I thank Jeff Walters, another committee member, for the resources and relationships he established at Eglin Air Force Base. I also thank my other committee members, Mike Aust, Jim Fraser, and Jim Parkhurst for their help. My research diverged greatly from the realm of Mike’s expertise, but he stuck with me and provided a fresh approach to my project. I also thank him for proving there is normalcy in academia and serving as a model of how a professor should be. Jim Fraser provided helpful direction with ecological and evolutionary aspects of my project. I thank Jim Parkhurst for his attention to detail. Through his tireless editing, I believe I have become a better writer. I thank the natural resources personnel at Eglin Air Force Base, especially Bruce Hagedorn and Dennis Teague. They were supportive of all aspects of my research and were understanding when things did not go as planned. I also thank James Furman and Kevin Hiers for their contributions to my project, friendship, guitar sessions, and introducing me to prescribed fire. I thank Jeff Herod and Theresa Thom for their iv expertise on stream ecology and collection of fish used in the tadpole experiment. I also thank Steve Laine and Kathryn Smith for their help with Eglin’s GIS data. I thank all the technicians who assisted me on some aspect of this project: Robert Emerson, Kathy Gault, Patrick Gault, Jim Kowalsky, Lourdes Oztolaza, Lou Phillips, and Renee Ripley. Their enthusiasm and assistance made this project succeed. I thank Dick Snyder and John Henry at the University of West Florida for access to their equipment and assistance with water analysis. I wish that aspect of my project would have worked. I thank Scott Klopfer, from the Conservation Management Institute, for his assistance with GIS analysis on Ch. 2. I thank Paul Moler, John Palis, and David Printiss for their input on flatwoods salamanders and Florida bog frogs. I also thank all the other biologists, who are too numerous to list, within the Department of Defense, Florida Natural Areas Inventory, Florida Fish and Wildlife Conservation Commission, Fish and Wildlife Service, Forest Service, and Nature Conservancy for contributing data, time, and ideas to this project in one form or another. I thank the Department of Fisheries and Wildlife Sciences at Virginia Tech for my education, both formally and informally. I thank the department, the College of Natural Resources, the Graduate School, and Dwight Chamberlain for additional financial support. Thanks to all the graduate students for making my four years at Tech very enjoyable. Thanks to my family, who always support my endeavors even though they don’t really understand what I do. Finally, thank you Mary for making all this worth while. v TABLE OF CONTENTS Abstract……………………………….…………………………………………….. ii Acknowledgements………………….………………………………………….…... iv Table of Contents…………………………………..…………………………..…… vi List of Tables.………....…….…………………….………………………………… viii List of Figures.…………………….………………………………….……………... x Foreword.……………….…………………………………………………….…….. xiii Chapter 1. Breeding biology, population dynamics, body size, and microhabitat of Florida bog frogs..…………….………..……………………….... 1 Methods…………………………………………………………………....……... 2 Results.…………………………………………………………….……………... 6 Discussion ………………………………………………………………….......... 8 Literature Cited.…………………………………….……………………………. 15 Tables.………………………………………………………….………………… 20 Figures………………………………………………………………….………... 24 Chapter 2. Movements, home ranges, and spatial interactions of Florida bog frogs and bronze frogs……………………………………………………………... 29 Methods…………………………………………………………………....…….. 61 Results.…………………………………………………………….……………... 33 Discussion ………………………………………………………………….......... 35 Literature Cited.…………………………………….……………………………. 42 Tables.………………………………………………………….………………... 46 Figures………………………………………………………………….………... 50 Chapter 3. Vocal repertoire of the Florida bog frog and response to conspecific and heterospecific calls……………………………………………………………... 58 Methods…………………………………………………………………....…….. 61 Results.…………………………………………………………….……………... 63 Discussion ………………………………………………………………….......... 66 Literature Cited.…………………………………….……………………………. 73 Tables.………………………………………………………….………………… 78 Figures………………………………………………………………….………... 79 vi Chapter 4. Individual voice recognition in Florida bog frogs…………………... 87 Methods…………………………………………………………………....…….. 89 Results.…………………………………………………………….……………... 91 Discussion …………………………………………………………………......... 92 Literature Cited.…………………………………….……………………………. 96 Tables.………………………………………………………….………………… 100 Figures………………………………………………………………….………... 103 Chapter 5. Response of Rana okaloosae, R. clamitans, and R. sphenocoephala tadpoles to chemical cues of snake and fish predators………………………….. 109 Methods…………………………………………………………………....…….. 111 Results.…………………………………………………………….……………... 113 Discussion ………………………………………………………………….......... 114 Literature Cited.…………………………………….……………………………. 117 Tables.………………………………………………………….………………… 121 Figures………………………………………………………………….……….... 123 Chapter 6. Burning trends and potential negative effects of suppressing wetland fires on flatwoods salamanders…………………………………………... 125 Methods…………………………………………………………………....……... 127 Results.…………………………………………………………….……………... 130 Discussion …………………………………………………………………........... 131 Literature Cited.…………………………………….…………………………….. 137 Tables.………………………………………………………….……………….... 144 Figures………………………………………………………………….……….... 148 Summary…………………………………………………………………………...... 151 Species profile: Florida bog frog………………………………………………..... 151 Management and research recommendations…………………………………….. 156 I. Florida bog frog………………………………………………………………… 156 II. Flatwoods salamander…………………………………………………………. 161 Literature Cited…………………………………………………………………… 165 Appendix…………………………………………………………………………..... 168 Vita………………………………………………………………………………….. 175 vii LIST OF TABLES Chapter 1. Breeding biology, population dynamics, body size, and microhabitat of Florida bog frogs Table 1.